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       Áø¼¼³ë»çÀ̵å Àü±¸Ã¼ ÇÕ¼º»ý¹°ÇÐ ¿¬±¸ µ¿Çâ ¹× Ãß¼¼ º¸°í¼­(¾ÆÁÖ´ë ÃÖº¸Çö, ÀÌÆòõ)
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       ÀÌÆòõ        2013.03.06 15:02        16457
´Ù¿î·Îµå : Áø¼¼³ë»çÀÌµå ¿¬±¸µ¿Çâ º¸°í¼­_ÀÌÆòõ.pdf(242 Kb)
 

Áø¼¼³ë»çÀ̵å Àü±¸Ã¼ ÇÕ¼º»ý¹°ÇÐ ¿¬±¸ µ¿Çâ ¹× Ãß¼¼ º¸°í¼­

 

ÃÖº¸Çö, ÀÌÆòõ

                                                                                                         ¾ÆÁÖ´ëÇб³

 

1. °³¿ä

 

Àü ¼¼°èÀûÀ¸·Î ±âÈĺ¯È­³ª ȯ°æ¿¡ ¿µÇâÀ» ÁÖÁö ¾Ê´Â ¿¡³ÊÁö¿Í ÇÕ¼º È­ÇÐ ¹°ÁúÀÇ °ø±Þ¿ä±¸¸¦ ´ëüÇÒ ¼ö ÀÖ´Â ÇÕ¼º »ý¹°ÇÐÀ» ÀÌ¿ëÇÑ Àç»ç¿ë ¿¡³ÊÁö¿Í È­Çй°ÁúÀÇ ¹ßÀüÀÌ ÀÌ·ç¾îÁö°í ÀÖ´Ù.¾ÆÀ̼ÒÇÁ·¹³ëÀ̵å (Isoprenoids)´Â ½ÄÇ°, ÀÇÇÐ, ¾àÇÐ, ¹ÙÀÌ¿À ¿¬·á ¹× È­ÇÐ µî ´Ù¾çÇÑ ºÐ¾ßÀÇ »ê¾÷ü¿¡ Àû¿ëµÇ°í ÀÖ´Â À¯±â ÇÕ¼ºÃ¼ÀÇ Å« ±×·ìÀÌ´Ù. ÀÌ·¯ÇÑ ´Ù¾çÇÑ ±¸Á¶ÀÇ ¾ÆÀ̼ÒÇÁ·¹³ëÀ̵å´Â »ýÇÕ¼º Àü±¸Ã¼ÀÎ Isopentenyl diphosphate (IPP)¿Í À̼ºÁúüÀÎ dimethylallyl diphosphate (DMAPP)·ÎºÎÅÍ ÇÕ¼ºµÇ¸ç, ÀÌ·¯ÇÑ Àü±¸Ã¼´Â mevalonate (MVA) pathway¿Í 2-C-methly-D-erythritol 4-diphosphate (MEP) pathway µÎ °³ÀÇ µ¶¸³ÀûÀÎ »ýÇÕ¼º ´ë»çȸ·Î¸¦ ÅëÇØ »ý»êµÈ´Ù.(±×¸² 1.)

 

Isoprenoids °è¿­ ¹°ÁúµéÀº º¸Åë ½Ä¹°Ã¼·ÎºÎÅÍ ºÐ¸®µÇ¸ç À̸¦ ÀÌ¿ëÇØ anticancer drugs, antimalarial drugs, »ó¾÷Àû ¿ëµµÀÇ flavor, fragranceµî¿¡ »ç¿ëµÇ°í ÀÖ´Ù. ±×·¯³ª ½Ä¹°Ã¼·ÎºÎÅÍÀÇ Á¶Á÷ ÃßÃâÀ» ÅëÇÑ ºÐ¸®´Â ÀϹÝÀûÀ¸·Î ³·Àº ¼öÀ²ÀÇ ¾çÀ» ¾òÀ» ¼ö ÀÖ¾î microbial host¸¦ ÀÌ¿ëÇÑ ³ôÀº ¼öÀ²ÀÇ product¸¦ ¾ò´Â ¿¬±¸ÀÇ º»°ÝÈ­°¡ Àý½ÇÇÑ ½ÃÁ¡ÀÌ´Ù. º» º¸°í¼­¿¡¼­´Â ÀÌ·± Isoprenoids Àü±¸Ã¼ÀÇ ÇÕ¼º»ý¹°ÇÐÀû ¿¬±¸ µ¿ÇâÀ» ¾Ë¾Æº¸°í À̸¦ ÀÌ¿ëÇÑ Çö½ÃÁ¡¿¡¼­ÀÇ ¹ßÀü ¹æÇ⼺À» °íÂûÇÏ°íÀÚ ÇÑ´Ù.

±×¸² 1. IsoprenoidsÀÇ Àü±¸Ã¼ »ýÇÕ¼ºÈ¸·Î: MEP ´ë»çȸ·Î¿Í MVA ´ë»çȸ·Î

(Jonathan Lombard & David Moreira, 2011)

 

2. Isoprenoids Àü±¸Ã¼ ´ë»çȸ·Î À¯ÀüÀÚÀÇ ´Ù¾ç¼º

 

ÀϹÝÀûÀ¸·Î Listeria monocytogenes ¿Í ƯÁ¤ streptomyces Á¾°ú °°ÀÌ µÎ °³ÀÇ IPP »ýÇÕ¼º ´ë»çȸ·Î¸¦ ¸ðµÎ °¡Áø °æ¿ì(Boucher & Doolittle et al, 20000)¸¦ Á¦¿ÜÇÏ°í´Â µ¿¹°°è¿¡¼­ IPP »ýÇÕ¼º °úÁ¤Àº MVA ´ë»çȸ·Î¿¡, °í¼¼±ÕÀº ¾à°£ÀÇ º¯ÇüµÈ MVA´ë»çȸ·Î¿¡ ƯȭµÇ¾î ÀÖÀ¸¸ç ´ëºÎºÐÀÇ ¹ÚÅ׸®¾Æ Á¾ÀÇ °æ¿ì µÎ °³Áß ÇϳªÀÇ ´ë»çȸ·Î°¡ µ¶Á¡ÀûÀ¸·Î Á¸Àç ÇÑ´Ù. MVA ´ë»çȸ·Î¿¡´Â ÃÑ 6°³ÀÇ È¿¼Ò°¡ °ü¿©Çϸç ÀÌ È¿¼ÒµéÀº acetyl-CoA ¿Í acetoacetyl-CoA¸¦ 3-hydroxy-3-methylglutaryl-CoA (HMG-CoA)·Î ÀüȯÇϸç Áß°£ ´ë»çüÀÎ mevalonate¿¡ phosphorylation °úÁ¤À» ÅëÇØ phosphomevalonate¿Í diphosphomevalonate¸¦ ÇÕ¼ºÇϸç ÃÖÁ¾ÀûÀ¸·Î C5 building blockÀÎ IPP³ª DMAPP·Î Àüȯ½ÃŲ´Ù.(±×¸² 1.)

 

MVP¿¡ ºñÇØ MEP ´ë»çȸ·ÎÀÇ °æ¿ì 8 ´Ü°èÀÇ È¿¼Ò È°¼º °úÁ¤À» ÇÊ¿ä·Î Çϸç pyruvate¿Í glyceraldehyde 3-phosphoate¸¦ 1-deoxy-D-xylulose 5-phosphate (DOXP)·Î ÇÕ¼ºÇϴ ù ¹ø° ´Ü°è¸¦ °ÅÃÄ DXR¿¡ ÀÇÇØ 2C-methyl-D-erythritol 4-phosphate (MEP)·Î ÀüȯµÈ´Ù. 4-diphosphocytidyl-2c-methyl-D-erythritol°¡ IspDÀÇ Ã˸ŷΠÀÎÇØ CTP¿Í MEP ¹ÝÀÀÀ¸·Î ÇÕ¼ºµÈ ÈÄ IspE¿¡ ÀÇÇØ CDP-ME2P·Î ÇÕ¼ºµÇ°í ¸¶Áö¸·À¸·Î IspF, IspG, IspH¿¡ ÀÇÇØ IPP·Î ÇÕ¼ºµÈ´Ù. (±×¸² 1.)

 

ÀÌ·¯ÇÑ Isoprenoids »ýÇÕ¼º Àü±¸Ã¼ÀÎ IPPÀÇ »ýÇÕ¼º ´ë»çȸ·Î¿¡ ´ëÇÑ ¿¬±¸´Â ´Ù¾çÇÑ ¹Ì»ý¹°¿¡ ´ëÇÏ¿© ÁýÁßÀûÀ¸·Î ÁøÇàµÇ¾î ¿Ô´Ù. ÃÖ±Ù¿¡´Â phylogenetic ¿¬±¸¸¦ ÅëÇÑ MVA ´ë»çȸ·Î À¯ÀüÀÚµéÀÇ À¯·¡ ¹× Á¾·ù¿¡ ´ëÇÑ ¿¬±¸°¡ È°¹ßÈ÷ ÁøÇàµÇ°í ÀÖÀ¸¸ç À̸¦ ÅëÇØ MEP ´ë»çȸ·Î¿Í MVA ´ë»çȸ·ÎÀÇ ÁøÈ­·ÐÀû ¿¬±¸°¡ ÁøÇà ÁßÀ̸ç, ´Ù¾çÇÑ ¹Ì»ý¹° Á¾µéÀÇ MVA ´ë»çȸ·Î À¯ÀüÀÚ ½Ã½ºÅÛÀÌ ¹àÇôÁö°í ÀÖ´Â Ãß¼¼ÀÌ´Ù.( Ç¥ 1.)

 

Ç¥ 1. Archaeal Genera¿Í Bacteria¿¡ °ü°èµÈ MVA ´ë»çȸ·Î À¯ÀüÀÚµéÀÇ Á¾º° ºñ±³

(Jonathan Lombard & David Moreira, 2011)

 

3. Áø¼¼³ë»çÀ̵å Àü±¸Ã¼ ´ë»çȸ·ÎÀÇ ÁøÈ­·ÐÀû ¹æÇ⼺

 

»ý¹°°èÀÇ ¼¼ °³ÀÇ Å« µµ¸ÞÀÎ (¿øÇÙ»ý¹°°è, ÁøÇÙ»ý¹°°è, °í»ý¹°°è)¿¡¼­ Áø¼¼³ë»çÀ̵å Àü±¸Ã¼ÀÎ IPPÀÇ »ýÇÕ¼º ´ë»ç °æ·Î´Â ¸Å¿ì º¹ÀâÇÏÁö¸¸ MEP ´ë»çȸ·Î´Â ¹ÚÅ׸®¾Æ¿¡¼­, ÀϹÝÀûÀÎ MVA ´ë»çȸ·Î´Â ÁøÇÙ»ý¹°°è, º¯ÇüµÈ MVA ´ë»çȸ·Î´Â °í»ý¹°°è¶ó´Â ¸íÈ®ÇÑ °ü°è°¡ ÀÖ´Ù. phylogenetic ¿¬±¸¸¦ ÅëÇÑ °¡¼³À» º¸¸é MVA ¿Í MEP ´ë»çȸ·ÎÀÇ ÁøÈ­·ÐÀû ½Ã³ª¸®¿À 3°¡Áö °¡´É¼ºÀÌ ÀÖ´Ù´Â ÁÖÀåÀÌ ÀÖ´Ù.(Lombard and Moreira, 2010). ù ¹ø° °¡¼³Àº Isoprenoids Àü±¸Ã¼ ´ë»çȸ·Î°¡ Á¸ÀçÇÏÁö ¾Ê´ø Ãʱâ Á¶»óÀ¸·ÎºÎÅÍ 3°³ÀÇ µµ¸ÞÀÎÀ¸·Î ÁøÈ­°úÁ¤¿¡¼­ °¢°¢ÀÇ Àü±¸Ã¼ ´ë»çȸ·Î°¡ ³ªÅ¸³µ´Ù´Â ¼³(±×¸² 2.A)À̸ç, µÎ ¹ø° °¡¼³Àº Ãʱâ Á¶»ó¿¡ MEP, MVA, MVA-Arc ´ë»çȸ·Î°¡ ¸ðµÎ Á¸ÀçÇÑ »óÅ¿¡¼­ 3°³ µµ¸ÞÀÎÀ¸·Î ÁøÈ­°úÁ¤¿¡¼­ °¢ ´ë»çȸ·Î¿¡ ¸Â°Ô ƯȭµÇ¾ú´Ù´Â ¼³ÀÌ´Ù(±×¸² 2.B ). ¸¶Áö¸· ¼¼ ¹ø° °¡¼³Àº Ãʱâ Á¶»ó¿¡¼­ MVA ´ë»çȸ·Î¸¦ °¡Áø »óÅ¿¡¼­ 3°³ µµ¸ÞÀÎÀ¸·Î ÁøÈ­ °úÁ¤¿¡¼­ °¢ ´ë»çȸ·Î¿¡ ¸Â°Ô º¯ÀÌ°¡ ÀϾ´Ù´Â ¼³ÀÌ´Ù(±×¸² 2.C).

±×¸² 2. »ý¹°ÇÐÀû ¼¼ °³ µµ¸ÞÀο¡¼­ÀÇ ´Ù¸¥ Isoprenoid »ýÇÕ¼º ´ë»çȸ·ÎÀÇ ¿¹»ó ÁøÈ­µµ.

                                      MEP ´ë»çȸ·Î´Â ÆĶõ»ö, ÀÏ¹Ý MVA ´ë»çȸ·Î´Â ÃÊ·Ï»ö, º¯Çü MVA ´ë»çȸ·Î´Â

                                      ºÓÀº»öÀ¸·Î Ç¥½ÃµÇ¾úÀ½. º°¸ð¾çÀº °¢ ´ë»çȸ·ÎÀÇ ±â¿øÀ» ³ªÅ¸³»¾úÀ½.

 

4. ÇÕ¼º»ý¹°ÇÐÀû Àû¿ë ±â¼ú µ¿Çâ

 

Ãֱ٠ȯ°æ¿¡ ¹ÌÄ¡´Â ¿µÇâÀÌ Å©´ø ±âÁ¸ÀÇ ÇÕ¼º ±â¼úÀÇ º¸¿ÏÀ» À§ÇØ Ä£È¯°æ ±â¼úÀÇ ¹ßÀüÀÌ Àå·ÁµÇ´Â Ãß¼¼¿¡¼­ ÇÕ¼º»ý¹°ÇÐÀ» ÅëÇÑ ¿©·¯ ¿¬±¸°¡ ÀÌ·ç¾îÁö°í ÀÖ´Ù. Áø¼¼³ë»çÀ̵带 Æ÷ÇÔÇÑ °í±â´É¼º Isoprenoids¿¡ ´ëÇÑ °¡Ä¡ÀÇ Áõ°¡·Î ÀÇÇÐ, ¾àÇÐ ºÐ¾ß¿¡¼­ ÁÖ·Î ¿¬±¸°¡ ÀÌ·ç¾îÁö°í ÀÖÀ¸¸ç »ê¾÷È­¸¦ À§ÇÑ »ý»ê·®/¼öÀ² Áõ´ëÀÇ ¸ñÀûÀ¸·Î Àü±¸Ã¼ÀÎ IPP ´ë»çȸ·ÎÀÇ ¿¬±¸ ¶ÇÇÑ ¹üÀ§°¡ È®´ëµÇ¾î°¡´Â Ãß¼¼ÀÌ´Ù.

 

MEP ´ë»çȸ·ÎÀÇ Àüü È帧¿¡ ´ëÇÑ ¿ÏÀüÇÑ Á¤º¸°¡ ÃÖ±Ù¿¡ ¹àÇôÁø ÀÌ·¡·Î MEP ´ë»çȸ·Î¸¦ ÅëÇÑ ¿¬±¸°¡ È°¹ßÈ÷ ÁøÇàµÇ°í ÀÖÀ¸¸ç ÁÖ·Î ÀÇÇÐÀû ºÐ¾ß¿¡¼­ MEP ´ë»çȸ·ÎÀÇ ÇÙ½ÉÀÌ µÇ´Â È¿¼ÒÀÇ ¹ßÇöÀ» ¾ïÁ¦ÇÏ´Â inhibitor¸¦ ÀÌ¿ëÇÑ Ç×¹ÚÅ׸®¾Æ ¹°ÁúÀÇ °³¹ß (Handa. S et al, 2013, Guobin Cai et al. 2013) ¹× MEP ´ë»çȸ·Î¿¡ °ü¿©ÇÏ´Â È¿¼Ò ÀÚü¿¡ mutation engineering ±â¹ýÀ» Àû¿ëÇÑ ÀçÁ¶ÇÕ È¿¼Ò¸¦ ÀÌ¿ëÇØ ÀÎü¿¡ ¼Õ»ó Á¶Á÷¿¡¼­ ¹ß»ýÇÏ´Â È¿°ú¿¡ µû¸¥ ¾àÇÐÀû ¿¬±¸ (Parul Gupta et al 2013) ¿ª½Ã ÁøÇàµÇ°í ÀÖ´Ù.

 

MVA ´ë»çȸ·ÎÀÇ °æ¿ì ÀÎü ³» sterol °èÅë¿¡ °ü¿©ÇÏ´Â Áß¿äÇÑ ´ë»çȸ·ÎÀ̱⠶§¹®¿¡ ÀÇÇÐÀû, ¾àÇÐÀû ºÐ¾ß¿¡¼­ ¸¹Àº ¿¬±¸°¡ ÁøÇàµÇ°í ÀÖÀ¸¸ç ÃÖ±Ù¿¡´Â »ê¾÷ÀûÀÎ ¿ëµµ·ÎÀÇ Àû¿ëÀ» À§ÇÑ ¿¬±¸ ¿ª½Ã È°¹ßÇÏ°Ô ÁøÇàµÇ°í ÀÖ´Ù. ÀÇÇÐÀû, ¾àÇÐÀû ºÐ¾ß¿¡¼­´Â MEP ´ë»çȸ·Î¿¡¼­ÀÇ ¿¬±¸¿Í ¸¶Âù°¡Áö·Î ´ë»ç°úÁ¤¿¡¼­ ÇÙ½É ¿ªÇÒÀ» ÇÏ´Â È¿¼ÒÀÇ È°¼ºÀ» ¾ïÁ¦ÇÏ´Â (ÁÖ·Î 3-hydroxy-3-methylglutaryl-CoA reductase) Simvastatin°ú °°Àº inhibitor¸¦ ÀÌ¿ëÇÑ cancer cellÀÇ apoptosis¸¦ À¯¹ß½ÃÅ°´Â ¿¬±¸ (Archana Gopalan et al, 2013)³ª IPP¿Í DMAPPÀÇ °ú¹ßÇöÀ» ÅëÇÑ À¯¿ë Isoprenoid ¹°ÁúÀÇ »ý»ê·®/¼öÀ² Áõ´ë¸¦ À§ÇÑ ¿¬±¸ ¶ÇÇÑ È°¹ßÈ÷ ÁøÇà ÁßÀÌ´Ù. ¶Ç´Â ´ë»çȸ·Î¿¡ Á¸ÀçÇÏ´Â ¿©·¯ À¯ÀüÀÚµé Áß ºÒÇÊ¿äÇÑ À¯ÀüÀÚÀÇ Á¦°Å¸¦ ÅëÇÑ ´ë»ç¹°ÁúÀÇ °ú¹ßÇö ½Ã½ºÅÛÀ» ±¸ÃàÇÏ´Â ¿¬±¸ (Bilge Özayd©¥n et al, 2013) µî ´Ù¾çÇÑ approach·Î Isoprenoid Àü±¸Ã¼ ´ë»çȸ·Î¿¡ ´ëÇÑ ÇÕ¼º»ý¹°ÇÐÀû ¿¬±¸°¡ ÀÌ·ç¾îÁö°í ÀÖ´Â Ãß¼¼ÀÌ´Ù.(Ç¥ 3.)

 

±×·¯³ª ÀÌ·¯ÇÑ IPP Àü±¸Ã¼ ´ë»çȸ·ÎÀÇ ±¤¹üÀ§ÇÑ Àû¿ë ¹üÀ§¿Í ÀáÀç·Â¿¡ ºñÇØ ¿¬±¸´Â ¾ÆÁ÷ ÃʱâÈ­ ´Ü°èÀ̸ç È°¼ºÈ­ ¶ÇÇÑ ¹Ì¹ÌÇÑ ½ÇÁ¤ÀÌ´Ù. µû¶ó¼­ °íºÎ°¡ ¹°ÁúÀÎ Áø¼¼³ë»çÀ̵带 ´ë·®»ý»êÇÒ ¼ö ÀÖµµ·Ï IPP Àü±¸Ã¼ ´ë»çȸ·Î¿¡ ´ëÇÑ ±âÁ¸ÀÇ ³·Àº ¼öÀ²ÀÇ Á¦¾àÀ» ¶Ù¾î³ÑÀ» ¼ö ÀÖ´Â IPP Àü±¸Ã¼ °ú¹ßÇö ½Ã½ºÅÛÀÇ ±¸ÃàÀÌ ÇÕ¼º»ý¹°ÇÐÀ» Åä´ë·Î ÀÌ·ç¾îÁ®¾ß ÇÑ´Ù.

 

Ç¥ 3. IPP »ý»ê MEP¿Í MVA ´ë»çȸ·Î¸¦ ÀÌ¿ëÇÑ ´Ù¾çÇÑ ÇÕ¼º»ý¹°ÇÐÀû Àû¿ë ¿¬±¸ 

 

5. Âü°íÀÚ·á

1. Ajikumar, P.K, Xiao, W.-H, Tyo, K.E.J, Wang, Y, Simeon, F, Leonard, E, Mucha, O, Phon, T.H, Pfeifer, B , Stephanopoulos, G., Isoprenoid pathway optimization for Taxol precursor overproduction in Escherichia coli, 2010, Science, 330, 70-74
2. Archana Gopalan, Weiping Yu, Bob G. Sanders, Kimberly Kline, Simvastatin inhibition of mevalonate pathway induces apoptosis in human breast cancer cells via activation of JNK/CHOP/DR5 signaling pathway, 2013, cancer letter, 329, 9-16
3. Begley, M., Bron, P. A., Heuston, S., Casey, P. G., Englert, N.,Wiesner, J., Jomaa, H., Gahan, C. G. M. & Hill, C., Analysis of the isoprenoid biosynthesis pathways in Listeria monocytogenes revealsa role for the alternative 2-C-methyl-D-erythritol 4-phosphate pathway in murine infection. 2008, Infect Immun 76, 5392–5401.
4. Boucher, Y. & Doolittle, W. F., The role of lateral gene transfer in the evolution of isoprenoid biosynthesis pathways. 2000, Mol Microbiol 37, 703-716.
5. Cornish, R. M., Roth, J. R. & Poulter, C. D., Lethal mutations in the isoprenoid pathway of Salmonella enterica. 2006, J Bacteriol 188, 1444–1450.
6. Eberl, M., Hintz, M., Reichenberg, A., Kollas, A. K., Wiesner, J. &Jomaa, H.. Microbial isoprenoid biosynthesis and human cd T cell activation. 2003, FEBS Lett 544, 4–10.
7. Gabrielsen, M., Rohdich, F., Eisenreich, W., Gra¡§wert, T., Hecht, S.,Bacher, A. & Hunter, W. N.. Biosynthesis of isoprenoids: a bifunctional IspDF enzyme from Campylobacter jejuni. 2004, Eur J Biochem 271, 3028–3035.
8. Gophna, U., Thompson, J. R., Boucher, Y. & Doolittle, W. F., Complex histories of genes encoding 3-hydroxy-3-methylglutarylCoenzymeA reductase. 2006, Mol Biol Evol 23, 168–178.
9. Hammond, R. K. & White, D. C., Carotenoid formation by Staphylococcus aureus. 1970, J Bacteriol 103, 191–198.
10. Handa, S.a, Ramamoorthy, D, Spradling, T.J. Guida, W.C, Adams, J.H, Bendinskas, K.G, Merkler, D.J, Production of recombinant 1-deoxy-d-xylulose-5-phosphate synthase from Plasmodium vivax in Escherichia coli, 2013, FEBS Open Bio, 3, 124-129
11. Jonathan Lombard and David Moreira, origins and Early Evolution of the Mevalonate Pathway of Isoprenoid Biosynthesis in the Three Domain of Life, 2011, Mol. Biol, Evol, 28(1), 87-99.
12. Laupitz, R., Hecht, S., Amslinger, S., Zepeck, F., Kaiser, J., Richter,G., Schramek, N., Steinbacher, S., Huber, R. & other authors., Biochemical characterization of Bacillus subtilis type II isopentenyl diphosphate isomerase, and phylogenetic distribution of isoprenoid biosynthesis pathways. 2004, Eur J Biochem 271, 2658–2669.
13. Parul Gupta, Aditya Vikram Agarwal, Nehal Akhtar, Rajender Singh Sangwan, Surya Pratap Singh, Prabodh Kumar Trivedi, Cloning and characterization of 2-C-methyl-d-erythritol-4-phosphate pathway genes for isoprenoid biosynthesis from Indian ginseng, Withania somnifera, 2013, Protoplasma, 250, 285-295
14. Pe¢¥rez-Gil, J., Bergua, M., Boronat, A. & Imperial, S., Cloning and functional characterization of an enzyme from Helicobacter pylori that catalyzes two steps of the methylerythritol phosphate pathway for isoprenoid biosynthesis. 2010,  Biochim Biophys Acta 1800, 919–928.
15. Puan, K. J., Wang, H., Dairi, T., Kuzuyama, T. & Morita, C. T., fldA is an essential gene required in the 2-C-methyl-D-erythritol 4-phosphate pathway for isoprenoid biosynthesis. 2005,  FEBS Lett 579, 3802–3806.
16. Rohdich, F., Wungsintaweekul, J., Fellermeier, M., Sagner, S., Herz, S., Kis, K., Eisenreich, W., Bacher, A. & Zenk, M. H., Cytidine 59-triphosphate-dependent biosynthesis of isoprenoids: YgbP protein of Escherichia coli catalyzes the formation of 4-diphosphocytidyl-2-Cmethylerythritol. 1999, Proc Natl Acad Sci U S A 96, 11758–11763.
17. Sangari, F. J., Pe¢¥rez-Gil, J., Carretero-Paulet, L., Garc©¥¢¥a-Lobo, J. M. & Rodr©¥¢¥guez-Concepcio¢¥ n, M. A new family of enzymes catalyzing the first committed step of the methylerythritol 4-phosphate (MEP) pathway for isoprenoid biosynthesis in bacteria. 2010, Proc Natl Acad Sci U S A 107, 14081–14086.
18. Sinead Heuston, Maire Begley, Cormac G.M. Gahan and Colin Hill, Isoprenoid Biosynthesis in Bacterial Pathogens, 2012, Microbiology 158, 1389-1401.
19. Takagi, M., Kaneda, K., Shimizu, T., Hayakawa, Y., Seto, H. & Kuzuyama, T. Bacillus subtilis ypgA gene is fni, a nonessential gene encoding type 2 isopentenyl diphosphate isomerase. 2004, Biosci Biotechnol Biochem 68, 132–137
20. Wilding, E. I., Kim, D.-Y., Bryant, A. P., Gwynn, M. N., Lunsford, R. D., McDevitt, D., Myers, J. E., Jr, Rosenberg, M., Sylvester, D. & other authors . Essentiality, expression, and characterization of the class II 3-hydroxy-3-methylglutaryl coenzyme A reductase of Staphylococcus aureus. 2000, J Bacteriol 182, 5147–5152.




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